Behavior of juvenile American lobsters, Homarus americanus, under predation risk

The influence of predation risk and food deprivation on the behavior and activity of juvenile American lobsters, Homarus americanus Milne Edwards, was examined in single and paired individuals in laboratory experiments performed during 1988 and in the winter of 1991/92. In the presence of a predator (the tautog Tautoga onitis Linnaeus) restrained behind a barrier, single lobsters significantly reduced the time spent feeding at night, consumed fewer mussels, and quickly brought them back to shelter. Single lobsters did not forage during the day in any treatment. If deprived of food for 60 h, they consumed more mussels and spent more time walking than recently fed (12-h food-deprived) lobsters. Paired lobsters did forage during the day in the presence of a predator. The smaller lobsters (subdominant) in the pairs foraged for a longer time in the presence than in the absence of a predator and significantly longer than single individuals. Shelter occupancy was significantly shorter in single, recently fed lobsters in the presence of a predator compared to time spent sheltering in its absence. Among food-deprived lobsters, paired individuals spent a significantly shorter time within the shelter than single lobsters in the absence of a predator. Larger (dominant) lobsters, however, spent more time than subdominant lobsters within the shelter during all periods of the day. Without a predator, paired lobsters spent significantly more time than single ones in shelter-related activities. Under predation risk, subdominant lobsters concentrated shelter-building time during the day and built a higher percent of alternative shelters than either single or dominant lobsters. In the absence of a predator, paired lobsters walked in the open area for a significantly longer time than single ones in the absence of a predator. This apparently was associated with fighting between dominant and subdominant lobsters and the attempts of the larger lobster to drive the smaller one from its shelter. During the day, lobsters fought for a significantly longer time in the presence than in the absence of a predator. When the tautog was not constrained, mortality rate was similar in both single and paired lobsters. Mortality rate among subdominant lobsters, however, was seven times higher than among dominant lobsters. We suggest that the risk of predation interferes with the ability of single juvenile lobsters to acquire and consume food. They appear to trade off energetic consideration against risk of predation when foraging away from the shelter. The introduction of a conspecific competitor to the system may further increase risk (of the subdominant) to the predator. Intraspecific interactions tend to increase the risk of predation to smaller lobsters but increase the survival rate among larger lobsters. Received: 6 February 1995 / Accepted: 2 September 1997     

Survival of communally reared larval and juvenile lobsters,Homarus americanus

Survival of individually reared larval and juvenile stage lobsters, Homarus americanus (Milne-Edwards), was significantly higher than in corresponding groups of communally reared individuals. Among communally reared lobsters, the mortality rate was highest in the second-stage larvae and then progressively decreased in the later stages. The relationship between survival and duration of molt period of each life-cycle stage indicates that asynchronous molting in the groups of communally reared lobsters is a contributing factor to the higher mortality rate. The molting and mortality curves of communally held lobsters reared from the first larval to first or second juvenile stage showed best cross correlation at 0- or 1-day time lag. The decreased mortality rate observed in the later larval and juvenile stages appears to have resulted from the establishment of new behavior patterns. Group interactions which are influenced by numerous extrinsic and intrinsic factors lead to higher mortality rate (cannibalism) among communally reared lobsters.     

The timing of chemical signaling with urine in dominance fights of male lobsters (Homarus americanus)

Chemicals carried in the urine are a rich source of information about the identity, sex, aggressive motivation, and other attributes of an animal. In agonistic interactions, animals should be careful about disclosing such information, since it can be used by receivers to the disadvantage of the sender. By adjusting the timing of urine release, the signaler may still influence the behavior of the receiver to its own benefit. Here we investigate the urine signaling of American lobsters (Homarus americanus), necessary for the maintenance of dominance, using a catheter technique with high temporal resolution. We hypothesize that urine release in lobster fights is not continuous but restricted to moments that may elicit a reaction in the receiver beneficial to the sender. We found that eventual winners signaled more readily in the initial phase of the fight. Eventual losers did not show such initial urination. This suggests that a confident lobster may use urine signals to influence the behavior of its opponent into giving up the fight at an early stage. For both winners and losers, urine release is linked to offensive behavior and increases as the intensity of agonistic behavior increases. The coupling of urine release to offensive behaviors appears to improve both the reliability and efficacy of the chemical signal. Releasing urine under the increasing risk of inflicting injury during the fight provides a handicap that may ensure reliability of the threat signal. Furthermore, the immediate coupling of urine components to offensive behavior may help to consolidate the receiver"s memory of the signaler"s individual scent and thus facilitate future recognition of the dominant animal. Received: 6 October 1999 / Accepted 28 August 2000     

Reproductive biology of the red alga Polysiphonia lanosa (Ceramiales) in the Bay of Fundy, Canada

The purpose of this investigation was to establish reproductive phenology of the dioecious marine floridean red alga Polysiphonia lanosa (L.) Tandy. Specifically, the distribution and frequencies of male and female plants, timing of their sexual maturity, fecundity and success of fertilization were investigated. The presence of cystocarps was taken as an indication of syngamy. Populations of P. lanosa in the Bay of Fundy reproduce in spring, summer and autumn. Both male and female filaments were present in equal abundance at all sampling locations, grew in close proximity, and occurred with equal abundance at all elevations above chart datum. During the growing season most of the mature plants contained reproductive organs with apparently functional gametangia containing gametes the structure and behavior of which conformed to those reported in the literature for other species of the genus. Male and female plants of P. lanosa matured synchronously during late May to early July at all collecting sites. The ratio of spermatia to carpogonia calculated for the sampled population was high, 3000–4800 spermatia:1 carpogonium. At the end of the growing season (late October) the fertilization success was at its maximum of 50 to 90% and at its minimum in July at 24%. The three free-living life-history phases (females, males and tetrasporophytes) were abundant and occurred with equal frequency, indicating that a complete life cycle was common at the studied location. Received: 10 December 1996 / Accepted: 16 January 1997     

Effects of environmental manipulation on the locomotor activity and agonistic behavior of cultured juvenile American lobsters,Homarus americanus

The locomotor activity patterns and agonistic encounters of cultured juvenile (IX–XIII stage) American lobsters, Homarus americanus (Milne-Edwards) held in 8 experimental environments were investigated to examine their relative contribution to aggressive level. Three variables: (1) lobsters individually or communally maintained, (2) shelters present or absent, and (3) open or closed seawater system were tested in a set of factorial experiments. Locomotor activity of lobsters held in a 12 h: 12 h light: dark regime was recorded for individually separated lobsters for the first 5 days and for communally held lobsters for the next 5 days of a 10-day experimental period. All lobsters maintained individually had similar levels of activity. When the same individuals were communally maintained, there appeared to be social inhibition of activity. In addition, communally held lobsters without shelter were twice as active as those provided with shelter. The activity levels of groups of communally held lobsters with shelter progressively decreased over the experimental period. All lobsters became somewhat entrained to the light: dark cycle and were nocturnally active. However, the degree of entrainment was strongly influenced (P<0.001) by the presence or absence of shelter and other lobsters. Individually maintained lobsters all showed good entrainment, although those provided with shelter were slightly better entrained. The presence of conspecifics desynchronized activity patterns when shelter was not provided. Aggressive level, as measured by the frequency of agonistic encounters per 15-min observation period and the number and types of displays per encounter, decreased over an 8-day period for groups of 3 lobsters in all experimental environments. Lobsters maintained in a closed system initially showed higher frequencies of agonistic encounters than those held in an open system. The observed decrease in aggressive level with time primarily resulted from decreased activity levels. When two lobsters met, the probability of an agonistic encounter was about 0.8 throughout the experimental period, but decreasing activity levels resulted in progressively fewer encounters. The results suggest several approaches for limiting aggression among communally maintained lobsters in culture systems by environmental manipulation.     

Fungi of the Bay of Fundy

During the spring and summer of 1979, 48 species of geofungi and several sterile, unidentified fungi were isolated from two off-shore stations on the Bay of Fundy, Canada. Isolations were made from subsurface water and surface slicks on 6 agar media including one containing hexadecane, and from subsurface water baited with hemp seeds. Numbers of fungi were 10 to 100 times higher than those reported for other offshore stations and similar to those reported for a saltmarsh outflow. Species isolated during a high freshwater flow largely corresponded to a typical soil flora. Species isolated during a reduced freshwater flow were either marine in character or noted for association with area littoral seaweeds. Three fungi (Cladosporium cladosporioides, Trichoderma viride, and Gliocladium rosium) were tested for growth on media of increasing salinity. Growth was measured by dry weight production, pH drop and glucose used in liquid culture. C. cladosporioides showed responses typical of a marine fungus, T. viride of a terrestrial fungus and G. roseum was intermediate. The results are discussed in relation to the role of geofungi in the sea.     

Movements of whale sharks (Rhincodon typus) tagged at Ningaloo Reef, Western Australia

Whale sharks (Rhincodon typus Smith) aggregate seasonally (March–June) to feed in coastal waters off Ningaloo Reef, Western Australia. Pop-up archival tags were attached to 19 individuals (total lengths 4.5–11.0 m) at this location in early May of 2003 and 2004 to examine their horizontal and vertical movements. The long-term movement patterns of six whale sharks were documented, all of which travelled northeast into the Indian Ocean after departing Ningaloo Reef. They used both inshore and offshore habitats and made extensive vertical movements, occasionally to a depth of at least 980 m. Frequent up-and-down movements, diel vertical migration, and crepuscular descents were evident in the depth records. The sharks experienced ambient temperatures ranging between 4.2 and 28.7°C and encountered gradients of up to 20.8°C on dives.     

Haemocyanin concentration of juvenile lobsters (Homarus gammarus) in relation to moulting cycle and feeding conditions

Groups of laboratory-reared, juvenile lobsters (Homarus gammarus, 40–50 g wet weight) were given different types of food (shrimp, mixed invertebrates and pellets, bivalves, starved) for a period of 22 wk. Haemocyanin concentration was measured weekly, moult stage and moultings noted. Haemocyanin concentration decreased rapidly in the starved group (from 0.8–0.6 to 0.2 mM). The group fed with bivalves showed a small decline with time, to 0.5–0.3 mM at the end of the period, indicating a deficient diet. The haemocyanin in the other groups did not change significantly during the experiment. Moultings decreased in frequency and the intermoult period was prolonged when the quality of food was decreased, and in the starved group moulting ceased completely after 1.5 months. Haemocyanin concentration was lowest immediately after moult and increased gradually through stages C and D. Lobsters that were starved or maintained on the bivalve diet did not reach the same premoult haemocyanin levels as those fed with shrimps and mixed food.     

Yolk utilization and hatching time in the Canadian lobster Homarus americanus

During the course of its embryonic development, the Canadian lobster Homarus americanus Milne-Edwards exhibits steady increases in water content (56.2 to 86.8%) and ash (5.8 to 21.2%), and a progressive decrease in energy content from 6636 to 4292 cal/g dry weight. Mean dry weight of a single egg is 965 g, equivalent to 6.4 cal; a freshly hatched egg. The lobster hatches about 1,500 larvae per night over a period of 4 to 5 days. Dry weight, ash and calorific contents of larvae hatched on different days show considerable variations. After larvae hatch on the first day, continuous salt absorption by eggs to hatch on subsequent days leads to a steady increase in ash content from 143 g/larva hatched on the first day to 255 g/larva hatched on the fourth day, and consequently, to an increase in dry weight from 854 to 956 g/larva. Metabolism of embryos (0.1 cal/day), which are yet to be hatched on subsequent days, depletes the calorific content per unit weight (from 4637 to 3837 cal/g dry weight) as well as per larva (from 3.98 to 3.67 cal).Dedicated to my colleague K.-H. Schumann (Biologische Anstalt Helgoland) who died on December 12, 1969 due to a diving accident.     

Tissue distribution and excretion of 99mTc-disofenin in three marine species: Pleuronectes americanus (winter flounder), Homarus americanus (lobster), and Mya arenaria (soft-shell clam)

To determine the pharmacokinetics of a small lipophilic molecule in vivo, the distribution and accumulation of ^99mTc-radiolabelled disofenin (diisopropylacetanilide iminodiacetic acid) were traced during 1991–1992 by scintigraphy and gamma well counting in winter flounder (Pleuronectes americanus collected from Boston Harbor and Long Island Sound in 1992), lobsters (Homarus americanus collected from Massachusetts Bay in 1991), and soft-shell clams (Mya arenaria purchased in 1991). The agent was distributed throughout the bodies of lobsters within 12 s, throughout flounder within 40 s, and throughout clams within 2 min. It was concentrated most strongly by the liver of flounder, which contained 61.2±7.8% of the injected dose within 1 h of injection, and by the lobster hepatopancreas. Accumulation also occurred in the flounder kidney, lobster antennal glands, and the kidney and pericardial glands of clams. The compound was rapidly excreted from the flounder liver into the gall bladder, and from the lobster hepatopancreas into the stomach. The data suggest its excretion from the lobster antennal glands and clam kidneys. The rate of clearance of disofenin from the body varied among species: 99±2.1% of the initial dose remained in flounder sampled 16 to 24 h after injection, compared to 80.5±7% remaining in the lobster after 15 h, and 87.4±5.9% remaining in clams after 27 h. The clearance rates in flounder and lobster are considered to be minimum values because of the lack of gut activity in unfed individuals. Overall, these in vivo tracer studies establish the utility of scintigraphy for assessing the uptake and excretion of a lipid soluble compound in different taxa, and may be applicable for evaluating disease and/or environmental effects on organ function in marine animals.     

Movements of farmed and wild Atlantic cod (Gadus morhua) released in Trinity Bay,Newfoundland

Fishery scientists and managers are investigating the feasibility of enhancing annual recruitment to the northern Atlantic cod (Gadus morhua L.) stock complex off Labrador and northeastern Newfoundland through the release of farmed fish back into the sea. Release of newly matured fish and adults with farm-advanced fecundity would increase the spawning biomass. Enhancement efforts might be measurably successful in major bays that are year-round habitats for cod. To determine if farmed cod would remain and spawn in Trinity Bay, 14 fish with surgically implanted transmitters were released in November 1992. Sonic tracking confirmed that farmed cod released on the western side of Trinity Bay overwintered within the bay, and integrated with wild cod approaching spawning condition in April 1993. Blood plasma antifreeze levels confirmed that these wild cod had overwintered inshore in subzero waters. A spawning aggregation was found in July 1993, providing evidence that northern cod reproduce in Newfoundland bays. These findings suggest that it may be possible to increase the number of cod spawning inshore through the release of farmed fish.     

A laboratory energy balance for the larvae and juveniles of the American lobster Homarus americanus

A laboratory energy budget was constructed for the larvae and juveniles of the American lobster Homarus americanus Milne-Edwards fed brine shrimp, Artemia saline L. Measured energy flows included ingestion, egestion, excretion of ammonia, routine and fed metabolism, growth, and production of exuvia. Digestion and assimilation were calculated and minimum ration of protein necessary to sustain larval lobsters was estimated. No change associated with metamorphosis was observed in rates of excretion, fed metabolism, and production of exuvia. Routine metabolism is not significantly higher for larvae than for juveniles. Growth changes from exponential in larvae to a slower increase in post-larvae. Consumption reflects changes in other variables. Changes in energy partitioning and energetic efficiencies associated with metamorphosis are largely due to change in rate of growth.     

Thermal histories of brooding lobsters, <Emphasis Type="Italic">Homarus americanus</Emphasis>, in the Gulf of Maine

Although it is widely accepted that migration by ovigerous lobsters (Homarus americanus Milne Edwards) optimizes thermal conditions for embryonic development, temperatures experienced by freely moving lobsters have never been measured. The precise thermal histories of 30 ovigerous lobsters at large in the Gulf of Maine were recorded to compare thermal conditions experienced during a brooding season. Analysis of both the temperature and movement data revealed a clear difference between lobsters smaller and larger than the size when 50% of individuals are mature (carapace length 93 mm). Although small and large lobsters ultimately experienced a similar number of degree-days above 3.4°C (952.8 for small and 983.6 for large) from 25 September 2002 until 27 July 2003, large lobsters and their embryos experienced less extreme and less variable temperatures. They were exposed to more gradual cooling in the fall and more gradual warming in the spring. These data, which are the first to document the seasonal temperatures experienced by ovigerous lobsters, suggest that migrations do not necessarily increase the number of degree-days experienced by developing embryos, but do reduce the variation in their thermal regime.     

Changes in nucleic acids over the molt cycle in relation to food availability and temperature in Homarus americanus postlarvae

Postlarval lobsters Homarus americanus Milne Edwards hatched from three females collected in 1989 from Block Island Sound, Rhode Island were reared individually in the laboratory under nine treatment combinations of temperature (15, 18 and 21°C) and feeding (starved, low ration and full ration). Total RNA, DNA (mgind.^-1), RNA:DNA ratios and molt stage were determined for individuals sampled at daily intervals. Postlarval lobsters had high resistance to starvation. A majority of the lobsters survive 12 d of food deprivation, with some surviving up to 24–29 d. During starvation, cell biomass (estimated from protein:DNA) decreased to a minumum size, whereas cell number (based on total DNA) was generally conserved. The molt cycle was arrested at molt stage C in the starved postalrvae. Instar duration was inversely related to temperature. However, the duration of the postlarval instar did not differ between the low and full ration treatments. Uncoupling of cell growth and the molt cycle was evident in the full and low ration treatments. In the full ration treatments, the postmolt through early premolt was characterized by a rapid increase in total RNA and DNA. Maximum cell biomass was attained by molt stage C or D₀. In the low ration treatments, total RNA and DNA were less than those in the full ration treatments and the maximum cell biomass was attained only at molt stage D_2-3. Notably, different feeding regimes resulted in different patterns in RNA:DNA ratios over the molt cycle during the postlarval instar. RNA:DNA ratio was a sensitive indicator of recent (2 to 3 d) food deprivation. However, this ratio was positively related to the level of feeding only at molt stages C to D₁ and was inversely related to temperature regime. These results indicate that the use of RNA:DNA ratios to estimate the relative nutritional state of postlarval tobsters must be qualified with respect to the period of the molt cycle and the temperature regime.     

Transmission of Pseudoterranova decipiens (Nematoda: Ascaridoidea) via benthic macrofauna to sympatric flatfishes (Hippoglossoides platessoides,Pleuronectes ferrugineus,P. americanus) on Sable Island Bank,Canada

Larval ascaridoid nematodes were surveyed in three flatfish species, Hippoglossoides platessoides, Pleuronectes ferrugineus, and P. americanus, from Sable Island Bank in February and June 1989. Pseudoterranova decipiens (sealworm) abundances differed significantly with both host species, the heaviest levels being found in H. platessoides, and host length, infections generally increasing with length, but not between February and June samples. Moderate numbers of larvae were found in P. ferrugineus while P. americanus were seldom infected. Two third-stage P. decipiens larvae (L₃), 3.2 and 7.0 mm in length, were recovered from the cephalothoraxes of two Mysis mixta in H. platessoides stomachs while five L₃'s (2.8 to 8.7 mm in length) were found among food (polychaetes, cumaceans, tanaids, amphipods) in the stomach of a P. ferrugineus. Levels of P. decipiens infection appear to be directly related to consumption of suprabenthic prey, which commonly occur in the diet of H. platessoides but are exploited less frequently by P. ferrugineus and rarely by P. americanus.     

Reproduction ofCalanus glacialis under ice in spring in southeastern Hudson Bay,Canada

In situ egg production rates ofCalanus glacialis Jaschnov females caught under the ice cover of southeastern Hudson Bay, Canada, between early April and mid-June 1986 were measured by direct observations and by an egg-ratio method in order to investigate the role of ice microalgae and under-ice phytoplankton production in copepod reproduction. Egg production commenced, although at low rates, in early May, about 3 wk after the onset of measurable concentrations of ice microalgae at the ice-water interface. Egg production increased to about 12 eggs female^–1 d^–1 in early June, after the ice cover began to melt. The time taken by immature adult females to reach maturity in the field was estimated to be ca. 3 wk (between late April and mid-May) by visual evaluation of gonad maturity in preserved females. In the laboratory, females caught on 10 April and maintained in saturating concentrations of the diatomThalassiosira weissflogii took only a few days to produce eggs at high rates, indicatingC. glacialis was food-limited during the ice algal bloom at the interface. We suggest that ice algal grazing changes the timing, relative to high food concentrations in the water column, of reproduction inC. glacialis by promoting oogenesis and oocyte maturation.Contribution to the programme of GIROQ (Groupe interuniversitaire de recherches océanographiques du Québec)Please send all reprint requests to Dr. Runge at his current address: Institut Maurice-Lamontagne, Ministère des Pêches et Océans, C. P. 1000, Mont-Joli, Québec G5H 3Z4, Canada     

Movements and food habits of striped bass (Morone saxatilis) in Delaware Bay (USA) salt marshes: comparison of a restored and a reference marsh

 There has been much recent interest in restoration of salt-marsh habitats to their natural structure and function. However, the criteria for success of such restorations are not well-defined. As part of a larger program to evaluate the restoration of a former salt-hay farm bordering Delaware Bay, New Jersey, USA, we monitored the response of a large predator, the striped bass Morone saxatilis, to the restoration. During June to October 1998 we compared tidal and diel movements and food habits of juvenile and adult striped bass (n = 82, 212 to 670 mm fork length) between a restored marsh and an adjacent reference marsh with similar physical characteristics (depth, salinity, temperature). Striped bass movements at both sites were characterized by ultrasonic tracking with small, surgically implanted tags (21 d rated battery-life). Striped bass (n = 23, 421 to 610 mm fork length) were tagged and released near the main creek mouths at both the restored (n = 14) and reference (n = 9) marshes. At both sites, striped bass tended to move up the main creek during ebb tide. At the restored site, ebb tide upstream-movements ranged from 0.1 to 3.5 km from the main creek mouth (mean = 1.2 km). During the upstream movement, the fish typically stopped every 200 to 300 m (presumably to feed) for 1 to 2 h. At the reference site, few of the tagged fish moved farther than 100 to 200 m upstream from the main channel mouth at ebb tide, perhaps in response to somewhat lower dissolved oxygen at this site. During flood tide, tagged fish at both sites moved out into Delaware Bay, where they remained within 200 to 500 m of the creek mouth. Striped bass were sampled with gill nets to determine additional aspects of habitat use and food habits. Striped bass in both marshes were much more abundant at creek mouths (catch per unit effort, CPUE = 1.17) than in the upper reaches of the creeks (CPUE = 0.13). In the creek mouths, CPUE was greater at the restored site (CPUE = 1.8) than at the reference site (CPUE = 0.5). At both sites, most fish (approx. 80%) were collected on the late ebb or early flood tides, i.e. around low tide, when prey were presumably concentrated at the creek mouths. Stomach contents of bass from both restored and reference marshes (n = 59, 212 to 670 mm fork length) revealed that striped bass were eating mostly blue crab (Callinectes sapidus), grass shrimp (Palaemonetes vulgaris), sand shrimp (Crangon septemspinosa), mummichog (Fundulus heteroclitus), and various unidentifiable fishes (probably anchovies, Anchoa mitchilli, and Atlantic silverside Menidia menidia). In conclusion, the restored marsh supported larger numbers of striped bass than the reference marsh, but there was little difference in the pattern of creek utilization or food habits at either site. Thus, the restored marsh appears to be functioning in a similar manner to the reference marsh for these large predators. Received: 28 June 1999 / Accepted: 1 August 2000     

Ecology and Conservation of the Endangered American Burying Beetle (Nicrophorus americanus)

Field studies were conducted on the western population of the endangered American burying beetle (Nicrophorus americanus) in Oklahoma and Arkansas to determine its habitat affinities. A common cause of declining populations is some specialized adaptation that makes it difficult to respond to a rapidly changing habitat. We evaluated the hypothesis that N. americanus is a habitat specialist in its search for food, preferring mature forests with deep, humic soils. This hypothesis was rejected. Based on comparisons of niche breadth among syntopic congeners and niche overlap, N. americanus is relatively generalized in its use of a range of habitats when searching for food. It is likely that the generalist nature and the endangered status of N. americanus both derive from the fact that it is the largest member of its guild. In comparison to smaller species, N. americanus breeds on larger carcasses, which are more unpredictable in space and time. It is likely, therefore, that N. americanus must search over a larger area and greater diversity of habitats than its smaller congeners.